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Does colony have a sub-family preference in raising queens?

I have been puzzling over whether colonies tend to favor larvae from one or more sub-families to raise queens from.

Assume ten drone mates, we have ten sub-families. If one has a distinct genetic resistance to an outside threat such as disease or parasite, it will have a higher survival rate. Thus, over time, its percentage of the worker population will increase, say to 20%.

But the queen still lays eggs from that family at a 10% rate. Is there any evidence or studies that shows the bees have any feedback method that increases the odds of the successful sub-family in raising queens?

Note that from my experience rearing queens, I would say no. They will accept larvae from any source. BUT most queen rearing is started under the emergency impulse, which is actually rare in nature. In a crisis, they may not be selective, and accept whatever larvae might save the hive.

Given the luxury of time, as in swarming or supercedure, perhaps they are more selective.

It is beyond the scope of a simple post, but this would help explain to me why drone laying is suppressed in a queenright hive. Darwin was extremely troubled by haplodiploidy and I am not convinced by Hamilton’s explanation.

Re: Does colony have a sub-family preference in raising queens?

From Dr. John Connor's Bee Sex Essentials, "efforts to show that super-sisters favor queens from their sub-family over half-sister queens have failed to show there is a clear and significant bias." However, he also notes research which has shown that bees from the same sub-family can recognize each other, and that division of labor within the hive often corresponds to these sub-families (patrilines).

Re: Does colony have a sub-family preference in raising queens?

Re: Does colony have a sub-family preference in raising queens?

MethowKraig - I'll shoot an email off to Doc (Dr. Lawrence John Connor) and ask about his bibliography for a starter reading list. My intuition says some patrilines will be more "popular" than others - and I'll bet the tendency starts early, favoring certain grubs when food is scarce, especially during supercedure response, etc., but I'd rather read a study and repeat an experiment, or design a new one and let the bees tell me the answer, any time!

I may send a few emails around the universities and Randy Oliver and others and see what studies are out there. Good question, good post! This is right at the core of rearing quality queens, especially for those of us who are running colony trait tests, mixing and concentrating semen from desirable drones, and getting into I.I. Thank you for starting this one!

Indypartridge - perhaps you could post a list from the bibliography of LJC's Bee Sex Essentials...?

Re: Does colony have a sub-family preference in raising queens?

Given a stable population of colonies (pre-human influence), a single queen will, on average, produce one successful daughter queen.

For ease of the math, assume an average of ten drones per typical colony, thus ten sub-families each colony.

First problem: 90% of genetic material contributed by drones is "dead-ended" at the colony because 9 out of ten sub-families do not produce a queen.

Also, note that each queen will, on average, have ten drone sons that contribute genetic material to other colonies. But only 10% of those drones will actually be lucky enough to have a daughter queen.

Second problem: Without going into the math of Hamilton's "relatedness index," if you multiply relatedness by the 10% success rate, the result is that drone laying workers would be a far more effective way of passing on genetic material. This method would also increase genetic diversity dramatically because the male side of the genome would be passed on. It would also be directly influenced by the sub-familiy's success rate. The more workers the more drones.

Third problem: Drone laying workers are the hive's last resort effort to pass on its genetics. They have tried supercedure. They have tried emergency rearing. The only thing left is drone laying workers. Given that the math says otherwise, there must be some extreme danger in this method of reproduction. What is the danger?

In summary, only when genetic material gets into the queen is it passed on in the lineage. Having a good drone mate or two and thus a successful colony is dead ended most of the time.

UNLESS, as I noted in my start of this thread, there is some sort of feed back mechanism to increase the success rate of queen rearing for the best sub-families.

If the workers do not exhibit a preference, perhaps it is the queen herself. The increase in the sub-family pheromone, resulting from more viable workers surviving, could influence her directly.

Re: Does colony have a sub-family preference in raising queens?

.... But only 10% of those drones will actually be lucky enough to have a daughter queen.

From evolution point of view, it is very high success rate!

Originally Posted by MethowKraig

...In summary, only when genetic material gets into the queen is it passed on in the lineage.

I am not sure I understood this. In my understanding, the drones determines the lineage because they are haploid and thus, "pure line". In queen, since she is diploid, Mendel's law would work efficiently "diluting" the lineage: only 25% will keep lineage in each generation (1/4 of 1/4 in 2nd and so on). Because many drones inseminate the queen, it is really difficult to keep lineage via queen. In this sense, drones are "lineage" keepers.

Very interesting topic. I wish to know more on this subject. Selection may be at the queen and/or nurse-bees level. It is easy to imagine that nurse-bees could have some preferences who to choose to rise. From another hand, at normal circumstances, it is a queen who lays the egg into queen cell. It is known (?) that queen may choose the semen to be used - particularly from africanized drones. I do not know if queen could segregate semen from all drones participated in insemination or most of the semen is mixed?

Re: Does colony have a sub-family preference in raising queens?

Darwin was extremely troubled by haplodiploidy and I am not convinced by Hamilton’s explanation.

I used the wrong word. Should be, "Darwin was troubled by eusocial behavior." Eusocial behavior means some of the females give up reproducing to help the queen.

However, it is interesting to note that Bees belong to the Hymenoptora Order, along with ants and wasps. The entire order reproduces by haplodiploidy (haploid males). There seems to be a link between haploidiploidy and social behavior. Almost all social insects are in the Hymenoptera Order. The one big exception is termites.

Re: Does colony have a sub-family preference in raising queens?

Sergey,

The queen is inseminated by many drones and thus her colony is influenced by many drones. However, only sub-family lines that actually produce a queen extend their drone's influence. If only one or two sub-families produce a queen, the other drone lines are at a dead end in the colony.

Re: Does colony have a sub-family preference in raising queens?

Originally Posted by MethowKraig

... If only one or two sub-families produce a queen, the other drone lines are at a dead end in the colony.

Agree. I think my wording was not good - I meant that two "pure" lineages (mother/father) presented only in the drones the queen produced- each drone in the hive inherited ether mother or father haploid set of genes. Each drones from worker bees (assuming random semen) would have one out of many genotype presented in the inseminated queen (plus mother's set). I agree that only one drone preserved the lineage via the queen, but it is happened in any diploid organisms.
The subject of this thread is much more interesting if bees could recognize and possibly select the beneficial (?) genotype (or phenotype?).
Sergey

Re: Does colony have a sub-family preference in raising queens?

I'm in the beginning of trying to design an/some experiment(s). Using the centrifuge, homogenize the semen of two drones at a time, one having a visible trait such as Cordovan or Chartreuse Eyes, and the other not having the chosen visible trait. Instrumentally inseminate one group of queens with the visible trait and one group of queens without. Put the resulting queens in observer hives and keep quantitative records of resulting bees and daughter queens. Watch for and record as many behaviors related to patrilines / sub-families as you can think of.

Re: Does colony have a sub-family preference in raising queens?

Sergey - a point of interest regarding if the semen is mixed. Instrumental insemination can utilize "homogenized" semen from multiple drones by putting the various semen samples into a centrifuge and spinning them all together, or semen directly collected from drones, unmixed. I doubt that nature mixes it as thoroughly as 20 G's in a centrifuge, but the question is a good one.

I do remember reading a study that said that several queens that had been instrumentally inseminated were observed going out on a mating flight anyways (after I.I.), and that many did not. Your question makes me wonder if laboratory mixing being different than "Mother Nature mixing" (and perhaps number of drone matings) has anything to do with an I.I. queen's behavior as contrasted to a naturally mated queen's behavior.

Either way, if Mama Queen has some kind of control over which husband's sperm she's laying, that would seem a surprise - but it is not out of the realm of possibility.

Re: Does colony have a sub-family preference in raising queens?

Originally Posted by MethowKraig

Third problem: Drone laying workers are the hive's last resort effort to pass on its genetics. They have tried supercedure. They have tried emergency rearing. The only thing left is drone laying workers. Given that the math says otherwise, there must be some extreme danger in this method of reproduction. What is the danger?

Let me propose a hypothesis for the danger in drone laying workers. I will call it the "arms race restriction."

If one sub family has 1% of its workers laying drones, the incentive will be for the second to have 2%, etc. Very quickly, within a dozen or so generations, the drone population will be dominated by one or two drone lines. This will feed back as a reduction in sex alleles and brood failure due to diploid drones.

The same would be true if a sub family is preferential to raising its own queens.

The success of the colony would seem to depend on the complete suppression of any competition between families. After all, by its very nature, competition would tear apart the very cooperation that makes a beehive successful.

However, this still leaves open the concept of the queen having an influence. Sergey noted there is evidence of queens being preferential to africanized drone semen. Perhaps queens have been influencing survival all along but it was just never noticed.

Re: Does colony have a sub-family preference in raising queens?

Originally Posted by MethowKraig

If one sub family has 1% of its workers laying drones, the incentive will be for the second to have 2%, etc. Very quickly, within a dozen or so generations, the drone population will be dominated by one or two drone lines. This will feed back as a reduction in sex alleles and brood failure due to diploid drones.

The same would be true if a sub family is preferential to raising its own queens.

Usually only 1 to 2 workers become laying workers. I wonder if 2% of a 60,000 bee colony (=1,200 Laying Workers) would tolerate each other laying eggs in the same colony? Would they fight? Wouldn't their pheromones suppress other workers from becoming LW's? I think even 1% is too high a number.

I think that the greater likelihood would be workers from a patriline preferring it's own brood, if they can indeed tell the difference. In the supercedure and swarming responses, this would make sense, because the workers choose to supercede a queen, even to the point of killing Mama once the VQ has mated and begun laying. Swarming response is likely a group decision - workers and possibly the queen, too, involving several factors. Swarming is probably a goal of a colony.

In the emergency response, they are more likely to choose any healthy grub the right age, or closest to it, and tend to try to make many queens at first, only continuing feeding the one or two that are best suited to becoming a queen after the second day. Emergency queens are often superceded quickly anyways, likely because they were fed worker jelly for a day or two before being switched back to royal jelly in the queenless emergency event, and thus became an intercaste queen/worker. This would tend to reduce the number of ovarioles in a "queen's" ovaries, causing her to lay fewer eggs over her lifetime, and possibly her rate per day.

Re: Does colony have a sub-family preference in raising queens?

I guess I am puzzling why drone laying workers are suppressed at all in terms of evolution since it would seem to be a better way to pass on genetics, at least by the math.

But if one family started producing lw's, couldn't it set off an arms race, where the result after a few generations would be one or two families dominating and reducing the number of sex alleles.

In other words, competition between families, in and of itself, would defeat the integrity of the colony. This would be true even if the competition was in the form of favoring their own larvae for starting a queen cell.

Competition between families thus might be very dangerous and the honeybee has evolved to suppress any urge that would threaten the cooperation between families. Sort of a bargain with the devil, "let's agree not to fight so we are stronger as a group." I realize that is an anthropomorphism.

The queen on the other hand, is sort of impartial. She might be responding to the pheromones in the colony. If one family is more successful, their numbers would be higher and she could then favor their sperm in producing queens.

Researchers have apparently looked to see if workers favor queen larvae from their own family and not found any evidence.

Maybe they just have never looked at the queen because it seems so impossible to imagine she can select sperm.

I'm not sure how you could design an experiment. If queens are selecting sperm, it might be hard to duplicate with I.I. because perhaps the sperm inside of her is actually in globules of mucus that keep the different lines separate. There is no way an instrument can duplicate what happens in flight.

I'm thinking this is a topic for an advanced lab to investigate. With genetic testing, I'm sure they would figure out a way.

Re: Does colony have a sub-family preference in raising queens?

MHK - I believe a healthy queen can lay more drones to pass male-passed traits along than LW's can, as the LW's time window before the hive dies out is about 44 days at most, minus 24 days to raise a drone, minus how many days it takes him to mature to mating ability. Also, a laying queen will lay plenty of worker grubs who will become nurse bees a few days later to help raise them and later feed them. These workers seem to know that drones are a luxury, and without a queen and no hope of making one, they are a last resort. I would probably wager that is why LW's laying drones is suppressed until the very end.

I doubt the math favors LW's over a healthy laying queen in evolution - the LW drones are often underfed and probably have a lower % chance of sucessfully mating and producing a colony that survives if conditions are less than ideal when the queenless / no viable grubs event occurred. Too late in the season - they'll lose a robbing fight and die off, etc. A queen that lives 4 years and makes one swarm plus a batch of healthy drones a year has done her tribe proud in terms of evolution.

Re: Does colony have a sub-family preference in raising queens?

Originally Posted by MethowKraig

Assume ten drone mates, we have ten sub-families. If one has a distinct genetic resistance to an outside threat such as disease or parasite, it will have a higher survival rate. Thus, over time, its percentage of the worker population will increase, say to 20%.

I think this intra-hive evolutionary force of sub-family attrition through natural selection due to stresses will have a lot to do with who gets to become queen over an acceptible evolutionary time period.

Re: Does colony have a sub-family preference in raising queens?

Charlie,

I think this thread got sidetracked on the LW issue. I agree with how you describe LW's as they are now. I was trying to understand why honeybees evolved that way.

But going back to the original topic. Given a hive with an exceptionally healthy sub family. In my example, growing from 10% to to 20% of the population through attrition of the other bees. It seems like there must be some mechanism for the hive to take advantage of this.

I can think of only two possibilities. Either could be controlled by pheromones.

1. Nurse bees influence the outcome of queen larvae by favoring the thriving family when raising queens.
2. The queen influences the outcome by favoring the family's sperm when laying in a queen cell.

Sounds like the first may have been studied but not the second.

I wish some experts would weigh in on this thread. Do you have any contacts with better minds than ours?

Re: Does colony have a sub-family preference in raising queens?

Dr. Larry Connor emailed me back, saying that there were conflicting results. He gave a brief list of study leads from the entomologists at UC Davis, Berkley, North Carolina State, and some names to look for - Harry Laidlaw, Robert Page, Radniks, others, David Tarpy, Otto Mackenson - and suggested that I ask Dr. Tom Rinderer at Baton Rouge Bee Lab. I have not yet heard back from Dr. Rinderer. It could be a while searching these studies without titles, but the topic is interesting.

My guess would be that hypothesis #1 is indeed worthy of investigating if it has not been done already, and that possibly in hypothesis #2 that Daddy Drone may have flavored his own sperm with whatever nectar / pollen beer his working ladies fed him, along with his mama's gene mix.

And Ojai, CA is in Ventura County (Just "North" err, West, really of LA) about 12 miles inland from the city of Ventura. I'm a short ways out in the sticks from Ojai.

Re: Does colony have a sub-family preference in raising queens?

The basis of Hamilton's thesis for the evolution of eusocial behavior is that a worker gives up her ability to reproduce because her own offspring would be related by 1/2 but helping her mother queen produces super-sisters that are related by 3/4.

Where Hamilton breaks down for me is she is also cooperating in raising half-sisters and drone brothers who are related only by 1/4.

This is noted in the above reference under: Current Therories: Haplodiploidy/Kin selection in the statement, "Other problems with this theory are when there are multiple males breeding with the queen then siblings are less related."

So, that is why I am wondering why workers wouldn't have evolved to raise their own drones, which would be related by 1/2. Obviously, over evolutionary time, they would have evolved to be well fed, etc. You could imagine a system where the workers laid haploid drones and the queen laid diploid workers.

There is some huge impediment to this happening. Again, I'm thinking there is some feedback mechanism, perhaps the sex allele, that stops this evolution.